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''Oreopithecus'' (from the Greek , and , , meaning "hill-ape") is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
genus of hominoid primate from the Miocene epoch whose fossils have been found in today's Tuscany and Sardinia in Italy. It existed nine to seven million years ago in the Tusco-Sardinian area when this region was an isolated island in a chain of islands stretching from central Europe to northern Africa in what was becoming the Mediterranean Sea. ''Oreopithecus'' was one of many European immigrants that settled this area in the VallesianTurolian transition and one of few hominoids, together with '' Sivapithecus'' in Asia, to survive the so-called Vallesian Crisis. To date, dozens of individuals have been discovered at the Tuscan localities of
Montebamboli Montebamboli is a hamlet and località in the comune of Massa Marittima, Tuscany, Italy. The settlement was first mentioned in a parchment from the year 754, and it is located in the hills of what is now the centre of Massa Marittima, within t ...
, Montemassi, Casteani, Ribolla, and, most notably, in the fossil-rich lignite mine in the Baccinello Basin, making it one of the best-represented fossil apes.


Evolutionary history

''Oreopithecus bambolii'' was first described by French paleontologist Paul Gervais in 1872, after the discovery of a juvenile mandible by Professor Igino Cocchi in a lignite mine at Montebamboli in 1862. In 1890, nearly a dozen new specimens were reported by Guiseppe Ristori, among them an upper jaw. In 1898, a left lower jaw was described by Felice Ottolenghi. In 1907, Giuseppe Merciai reported four maxillae and a lower jaw from the Grosseto mine at Ribolla. During this period there was no consensus whether ''Oreopithecus'' was a monkey or an ape. From 1949 onwards, Swiss paleontologist began to restudy the known material. In 1954, 1955, 1956 and 1958 he claimed ''Oreopithecus'' were a true hominin—based on its premolars, short jaws and reduced canines, at the time considered diagnostic of the hominin family. This hypothesis was immediately hotly discussed by his fellow palaeontologists. When he toured the world to give a series of lectures, his views generated an enormous press coverage, often being presented as a challenge to the Darwinian descent of man from apes. After Hürzeler was invited to give a lecture in New York in March 1956, the
Wenner-Gren Foundation Axel Lennart Wenner-Gren (5 June 1881 – 24 November 1961) was a Swedish entrepreneur and one of the wealthiest men in the world during the 1930s. Early life He was born on 5 June 1881 in Uddevalla, a town on the west coast of Sweden. He ...
decided to finance excavations in Italy, with the cooperation of the Italian paleontologist Alberto Carlo Blanc. On 2 August 1958, Hürzeler's views seemed to be confirmed when he discovered a complete skeleton in Baccinello, which in 1960 he interpreted as a biped because of the short
pelvis The pelvis (plural pelves or pelvises) is the lower part of the trunk, between the abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also called bony pelvis, or pelvic skeleton). The ...
was closer to those of hominins than those of
chimpanzee The chimpanzee (''Pan troglodytes''), also known as simply the chimp, is a species of great ape native to the forest and savannah of tropical Africa. It has four confirmed subspecies and a fifth proposed subspecies. When its close relative th ...
s and gorillas. Hominin affinities claimed for ''Oreopithecus'' remained controversial for decades until new analyses in the 1990s reasserted that ''Oreopithecus'' was directly related to '' Dryopithecus''. The peculiar cranial and dental features were explained as consequences of
insular Insular is an adjective used to describe: * An island * Someone who is isolated and parochial Insular may also refer to: Sub-national territories or regions * Insular Chile * Insular region of Colombia * Insular Ecuador, administratively known ...
isolation. This new evidence confirmed that ''Oreopithecus'' was bipedal but also revealed that its peculiar form of bipedalism was much different from that of '' Australopithecus''. The hallux formed a 100° angle with the other toes, which enabled the foot to act as a tripod in erect posture, but prevented ''Oreopithecus'' from developing a fast bipedal stride. When a land bridge broke the isolation of the Tusco-Sardinian area , large predators such as '' Machairodus'' and '' Metailurus'' were present among the new generation of European immigrants and ''Oreopithecus'' faced quick extinction together with other endemic genera.


Taxonomic classification

Known as the "enigmatic hominoid", ''Oreopithecus'' can dramatically rewrite the palaeontological map depending on whether it is a descendant of the European ape '' Dryopithecus'' or an African anthropoid. Some have suggested the unique locomotory behavior of ''Oreopithecus'' requires a revision of the current consensus on the timing of bipedality in human developmental history, but there is limited agreement on this point among paleontologists. Simons (1960) considered ''Oreopithecus'' closely related to the early Oligocene '' Apidium'', a small arboreal anthropoid that lived nearly 34 million years ago in Egypt. ''Oreopithecus'' shows strong links to modern apes in its postcranium and, in this respect, it is the most modern Miocene ape below the neck, with closest similarities to the postcranial elements of '' Dryopithecus'', but its dentition is adapted to a leafy diet and a close link is uncertain. Others claim it to be either the sister taxon to Cercopithecoidea or an even direct human ancestor, but it is usually placed in its own subfamily within
Hominidae The Hominidae (), whose members are known as the great apes or hominids (), are a taxonomic family of primates that includes eight extant species in four genera: '' Pongo'' (the Bornean, Sumatran and Tapanuli orangutan); ''Gorilla'' (the ea ...
. It could instead be added to the same subfamily as ''Dryopithecus'', perhaps as a distinct tribe (Oreopithecini). A cladistic analysis of ''Nyanzapithecus alesi'' recovers ''Oreopithecus'' as a member of the proconsulid subfamily
Nyanzapithecinae The Nyanzapithecinae or Nyanzapithecines are a subfamily of extinct Dendropithecidae as sister of '' Simiolus''. The group contains ''Rangwapithecus'', '' Turkanapithecus'', '' Rukwapithecus'', ''Oreopithecus'', and ''Nyanzapithecus''. In the fo ...
.


Physical characteristics

''Oreopithecus bambolii'' is estimated to have weighed . It possessed a relatively short snout, elevated nasal bones, small and globular neurocranium, vertical orbital plane, and
gracile Gracility is slenderness, the condition of being gracile, which means slender. It derives from the Latin adjective ''gracilis'' (masculine or feminine), or ''gracile'' ( neuter), which in either form means slender, and when transferred for examp ...
facial bones. The shearing crests on its molars suggest a diet specializing in plant leaves. The very robust lower face, with a large attachment surface for the masseter muscle and a
sagittal crest A sagittal crest is a ridge of bone running lengthwise along the midline of the top of the skull (at the sagittal suture) of many mammalian and reptilian skulls, among others. The presence of this ridge of bone indicates that there are exceptiona ...
for attachment of the temporal muscle, indicates a heavy masticatory apparatus. Its teeth were small relative to body size. The lack of a diastema (gap) between the second incisor and first
premolar The premolars, also called premolar teeth, or bicuspids, are transitional teeth located between the canine and molar teeth. In humans, there are two premolars per quadrant in the permanent set of teeth, making eight premolars total in the mouth ...
of the mandible indicates that ''Oreopithecus'' had canines of size comparable to the rest of its dentition. In many primates, small canines correlate with reduced inter-male competition for access to mates and less sexual dimorphism.


Positional behavior

Its habitat appears to have been
swamp A swamp is a forested wetland.Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. 497 p. Swamps are considered to be transition zones because both land and water play a role in ...
y, and not savanna or forest. The postcranial anatomy of ''Oreopithecus'' features adaptations for suspensory arborealism. Functional traits related to suspensory locomotion include its broad thorax, short trunk, high intermembral index, long and slender digits, and extensive mobility in virtually all joints. Its fingers and arms seem to show adaptations for climbing and swinging. Its foot has been described as chimp-like, but is different from those of extant primates. The habitual line of leverage of the primate foot is parallel to the third metatarsal bone. In ''Oreopithecus'', the lateral metatarsals are permanently abducted so that this line falls between the first and second metatarsals instead. Furthermore, the shape of the tarsus indicate loads on the foot were transmitted to the medial side of the foot instead of the lateral, like in other primates. The metatarsals are short and straight, but have a lateral orientation increase. Its foot proportions are close to the unusual proportions of ''Gorilla'' and ''Homo'' but are distinct from those found in specialized climbers. The lack of predators and the limitation of space and resources in ''Oreopithecus''′ insular environment favored a locomotor system optimized for low energy expenditure rather than speed and mobility. ''Oreopithecus'' has been claimed to exhibit features that are adaptations to upright walking, such as the presence of a
lumbar In tetrapod anatomy, lumbar is an adjective that means ''of or pertaining to the abdominal segment of the torso, between the diaphragm and the sacrum.'' The lumbar region is sometimes referred to as the lower spine, or as an area of the back i ...
curve, in distinction to otherwise similar species known from the same period. Since the fossils have been dated to about 8 million years ago, this would represent an unusually early appearance of upright posture. However, a reevaluation of the spine from a skeleton of ''Oreopithecus'' has led to the conclusion that it lacked adaptations for habitual bipedality


Semicircular canals

The semicircular canals of the inner ear serves as a sense organ for balance and controls the reflex for gaze stabilization. The inner ear has three canals on each side of the head, and each of the six canals encloses a membranous duct that forms an endolymph-filled circuit. Hair cells in the duct's
auditory ampulla Auditory means of or relating to the process of hearing: * Auditory system, the neurological structures and pathways of sound perception ** Auditory bulla, part of auditory system found in mammals other than primates ** Auditory nerve, also known ...
pick up endolymph disturbances caused by movement, which register as rotatory head movement. They respond to body sway of frequencies greater than 0.1 Hz and trigger the vestibulocollic (neck) reflex and vestibuloocular (eye) reflex to recover balance and gaze stability. The bony semicircular canals allow estimates of duct arc length and orientation with respect to the sagittal plane. Across species, the semicircular canals of agile animals have larger arcs than those of slower ones. For example, the rapid leaper '' Tarsius bancanus'' has semicircular canals much bigger than the slow-climbing '' Nycticebus coucang''. The semicircular canals of brachiating
gibbon Gibbons () are apes in the family Hylobatidae (). The family historically contained one genus, but now is split into four extant genera and 20 species. Gibbons live in subtropical and tropical rainforest from eastern Bangladesh to Northeast India ...
s are bigger than those of arboreal and terrestrial quadrupedal great apes. As a rule of thumb, arc size of the ducts ''decreases'' with body mass and consequently slower angular head motions. Arc size ''increases'' with greater agility and thus more rapid head motions. Modern humans have bigger arcs on their anterior and posterior canals, which reflect greater angular motion along the sagittal plane. The lateral canal has a smaller arc size, corresponding to reduced head movement from side to side. Allometric measurements on the bony labyrinth of BAC-208, a fragmentary cranium that preserves a complete, undeformed petrosal bone suggest that ''Oreopithecus'' moved with agility comparable to extant great apes. Its anterior and lateral semicircular canal sizes fall within the range for great apes. Its relatively large posterior arc implies that ''Oreopithecus'' was more proficient at stabilizing angular head motion along the sagittal plane.


Dexterity

''Oreopithecus'' had hominin-like hand proportions that allowed a firm, pad-to-pad precision grip. Features present in the hands of neither non-human-extant nor fossil apes include hand length, relative thumb length, a deep and large insertion for the flexor pollicis longus, and the shape of the carpometacarpal joint between the metacarpal bone of the index finger and the capitate bone. At the base of the second metacarpal bone, the facet for the capitate is oriented transversally, as in hominins. The capitate, on the other hand, lacks the waisting associated with apes and climbing, and still present in '' Australopithecus''. ''Oreopithecus'' share the specialised orientation at the carpometacarpal joint with '' A. afarenis'' and the marked groove for the flexor pollicis longus with '' A. africanus''. It is thus likely that the hand morphology of ''Oreopithecus'' is derived for apes and convergent for early hominins.


See also

* List of fossil sites * List of human evolution fossils ''(with images)''


Explanations


Notes


References

* * * * * * * * * * *


External links

* — A photo of an ''Oreopithecus bamboli'' jaw * {{Taxonbar, from=Q131700 Miocene primates of Europe Prehistoric apes Fossil taxa described in 1872 Prehistoric mammals of Europe Prehistoric primate genera